European early modern humans

European early modern humans (EEMH) is a term for the earliest populations of anatomically modern humans in Europe, during the Upper Paleolithic. It is taken to include fossils from throughout the Last Glacial Maximum (LGM), covering the period of about 48,000 to 15,000 years ago (48–15 ka), spanning the Bohunician, Aurignacian, Gravettian, Solutrean and Magdalenian periods.

Cro-Magnon 1 skull (1884 drawing)

EEMH formed a genetically isolated population for more than 20,000 years, between c. 37 and 14 ka, with significant Mesolithic admixture from the Near East and Caucasus beginning from around 14 ka (in some regions not before 12 ka).[1] The description as "modern" is used as contrasting with the "archaic" Homo heidelbergensis and Homo neanderthalensis, who lived within Europe during about 400 ka to 37 ka, and who with the arrival of EEMH became extinct or absorbed into their lineage.

The oldest known EEMH fossil remains, confidently dated to before 40 ka, were found at Riparo Mochi (Italy), Geissenklösterle (Germany) and Isturitz (France).[1][2] Other known remains of EEMH can be dated to before 40 ka with some certainty: those from Grotta del Cavallo in Italy, and from Kents Cavern in England, have been radiocarbon dated to 43±2 ka.[3][4] A number of other early fossils are dated close to or just after 40 ka, including fossils found in Romania (Peștera cu Oase, 39.5±2.5 ka) and Russia (Kostenki-14, 37.5±2.5 ka).[5] The Siberian Ust'-Ishim man, dated to 45 ka, was not geographically found in Europe, and indeed is not part of the "Western Eurasian" genetic lineage, but intermediate between the Western Eurasian and East Asian lineages.[1]

The EEMH lineage in the European Mesolithic is also known as "West European Hunter-Gatherer" (WHG). These mesolithic hunter-gatherers emerge after the end of the LGM c. 15 ka and are described as more gracile than the Upper Paleolithic Cro-Magnons.[note 1] The WHG lineage survives in contemporary Europeans, albeit only as a minor contribution overwhelmed by the later Neolithic and Bronze Age migrations.

The term EEMH is equivalent to Cro-Magnon Man, or "Cro-Magnons", a term derived from the Cro-Magnon rock shelter in southwestern France, where the first EEMH were found in 1868. Louis Lartet (1869) proposed Homo sapiens fossilis as the systematic name for "Cro-Magnon Man". W.K. Gregory (1921) proposed the subspecies name Homo sapiens cro-magnonensis. In literature published since the late 1990s, the term EEMH is generally preferred over the common name Cro-Magnon, which has no formal taxonomic status, as it refers neither to a species or subspecies nor to an archaeological phase or culture.[note 2] Use of "Cro-Magnon" is mostly restricted to times after the beginning of the Aurignacian proper, c. 37 to 35 ka.[7]

Chronology
Map of the distribution of the main pre-LGM Aurignacian sites.
AMH fossil remains in Europe and Africa dated between 55 kya and 25 kya (calibrated carbon dates, as of 2013), aligned with climate reconstruction (North Greenland Ice Core Project, D–O events (fluctuations) indicated).[8]
Further information: Ahmarian, Emiran, Haplogroup N (mtDNA), Haplogroup IJ, Neanderthal admixture, and Neanderthal extinction

While anatomically modern humans (AMH) may have been present in West Asia since before 250 ka,[9] modern non-Africans mainly descend from the main successful out of Africa expansion at around 65–55 ka. This movement was an offshoot of the rapid expansion within East Africa associated with mtDNA haplogroup L3.[10] EEMH are associated with mtDNA haplogroup N, also widespread in Central Asia,[11] and with Y-chromosomal haplogroup CF.

AMH are estimated to have marginally interbred with Neanderthals during about 65 to 47 ka, most likely in West Asia soon after leaving Africa.[12][13] Ancestral West Eurasians began to move into Europe beginning about 45 ka. Neanderthals became extinct shortly after this time, presumably being outcompeted or actively killed by the advancing EEMH. Admixture with Neanderthals appears to cease almost entirely after 45 ka, in spite of several millennia of continued co-existence of AMH and Neanderthals in Europe.[14]

There are two main hypotheses as to the route taken by the earliest AMH entering Europe, following the Danubian corridor or the Mediterranean coast along the Balkans. Support for either hypothesis relies on accurate dating of the earliest known fossils in the region. High-precision dating of the earliest Proto-Aurignacian sites in Europe, Riparo Mochi (Italy), Geissenklösterle (Germany), and Isturitz (France), have yielded dates of close to 42 ka, indicating that EEMH spread throughout Western Europe very rapidly.[2]

EEMH sites in Europe earlier than 37 ka are also termed Proto-Aurignacian.[15] The Aurignacian proper, the stage associated with the original Cro-Magnon find, appears to have developed within Europe. It lasts from 37 ka until about 28 ka.

The Gravettian is the European culture immediately preceding the LGM, about 28 to 22 ka, but the early Gravettian overlaps with the later Aurignacian, from as early as 33 ka.[16] During the LGM proper, beginning about 22 ka, there are two main refugia, the Solutrean in Southwestern Europe, and the Epi-Gravettian in Italy and Southeastern Europe.

With advancing deglaciation after about 17 ka, finds associated with the Magdalenian are transitional to the mesolithic hunter-gatherer (WHG) populations. The European Mesolithic is taken to begin after about 14 ka (Azilian).

Physical attributes

Cro-Magnons were anatomically modern, straight limbed and tall compared to the contemporaneous Neanderthals. They are thought to have stood on average 1.66 to 1.71 m (5 ft 5 in to 5 ft 7 in) tall. They differ from modern-day humans in having a more robust physique and a slightly larger cranial capacity.[17] The Cro-Magnons had fairly low skulls, with wide faces, robust mandibles, blunted chins, narrow noses,[18] and moderate to no prognathism.[19] A distinctive trait was the rectangular eye orbits, similar to those of modern Ainu people. Their vocal apparatus was like that of present-day humans and they could speak.[20] Their brain capacity was about 1,600 cc (98 cu in), which is larger than the average for all modern humans.

Mitochondrial DNA analysis places the early European population as sister group to the East Asian groups of the Upper Paleolithic, dating the divergence to some 50,000 years ago.[21]

Analysis of ancient DNA of EEMH and Mesolithic fossils suggests that alleles related to the light skin characteristic of modern Europeans (TYRP1 SLC24A5 and SLC45A2) were not yet present. They are thought to have arisen during the LGM, most likely in the Caucasus, and were spread to Europe in a "selective sweep" during the late Upper Paleolithic to early Mesolithic, between c. 19 to 11 ka.[22][23] The HERC2 variation for blue eyes first appears around 14 ka in Italy and the Caucasus.[24]

Behavior and culture
Aurignacian painting of cave lions from Chauvet Cave, Ardèche (museum replica)
Aurignacian bone flute from Geissenklösterle (Swabia)

The flint tools found in association with the remains at Cro-Magnon have associations with the Aurignacian culture that Lartet had identified a few years before he found (in 1868) the first skeletons. The Aurignacian differ from the earlier cultures in their finely worked bone or antler points and flint points made for hafting, in the production of Venus figurines and in cave painting.[25] They pierced bones, shells and teeth to make body ornaments. The figurines, cave-paintings, ornaments and the mysterious Venus figurines are a hallmark of Cro-Magnon culture, contrasting with the utilitarian culture of the Neanderthals.[26]

Like most early humans, the Cro-Magnons were primarily big-game hunters, killing mammoth, cave bears, horses, and reindeer.[27] They hunted with spears, javelins, and spear-throwers. (Archery had not yet been invented.) They would have been nomadic or semi-nomadic, following the annual migrations of their prey, and would also have eaten plant materials. In the village of Mezhirich in Ukraine, excavations have unearthed several huts built from mammoth bones (possibly representing semi-permanent hunting camps).[20][28]

Finds of spun, dyed, and knotted flax fibers among Cro-Magnon artifacts in Dzudzuana show that European early modern humans made cords for hafting stone tools, weaving baskets, or sewing garments.[29] Apart from the mammoth bone huts mentioned, they constructed shelters of rocks, clay, branches, and animal hide/fur. Manganese and iron oxides were used in rock paintings.

Assemblages and fossils
Site Region (country) Industry Age (ka) Description
Grotta del CavalloItalyProto-Aurignacian44In November 2011, tests conducted at the Oxford Radiocarbon Accelerator Unit in England on what were previously thought to be Neanderthal baby teeth, which had been unearthed in 1964 from the Grotta del Cavallo in Italy. These were identified as the oldest anatomically modern remains ever discovered in Europe, dating from between 43,000 and 45,000 years ago. No tools were associated with the find.
GeissenklösterleCentral Europe (Germany)Proto-Aurignacian42The bone flute of Geissenklösterle (Swabia) has been radiocarbon dated to 43–42 ka. The so-called adorant from the Geißenklösterle cave is somewhat younger (40–35 ka). Other paleolithic art of the early Aurignacian was found in the Swabian Alb area, including Venus of Hohle Fels and the Lion-man figurine (see Caves and Ice Age Art in the Swabian Jura)[30] No dated human remains are associated with these finds.
Kostenki-14Eastern Europe (Russia)Proto-Aurignacian38A male from Kostenki-14 (Markina Gora), dated 35–40 ka, was found to have close ancestry with both "Mal'ta boy" (24 ka) of central Siberia (Ancient North Eurasian) and to the later Mesolithic hunter-gatherers of Europe and western Siberia, as well as with a basal population ancestral to Early European Farmers.[5]
Peștera cu OaseSoutheastern Europe (Romania)Proto-Aurignacian38
Oase 2 skull

Peștera cu Oase ("Bones Cave") near the Iron Gates in Romania appears to be a cave bear den; the human remains may have been prey or carrion. No tools are associated with the finds. Oase 1 holotype is a robust mandible which combines a variety of archaic, derived early modern, and possibly Neanderthal features. The modern attributes place it close to EEMH among Late Pleistocene samples. The fossil is one of the few finds in Europe which could be directly dated and is at least 37,800 years old. Oase 2 (and fragments Oase 3), discovered in 2005, is the skull of a young male, again with mosaic features, some of which are paralleled in the Oase 1 mandible.[31]

Kent's CavernWestern Europe (UK)Proto-Aurignacian36The Kent's Cavern find is a prehistoric maxilla (upper jawbone) fragment was uncovered in the cavern during a 1927 excavation in Kents Cavern by the Torquay Natural History Society, and named Kent's Cavern 4. In 1989 the fossil was radiocarbon dated to be from 36,400–34,700 years ago, although various fauna fossils in older strata present on the site have been dated in 2011 to 44,200–41,500 years ago and indirectly associated with the human remains, the latest datings and conclusions in question being highly disputed.[4]
SungirEastern Europe (Russia)Aurignacian34
Sungir 2

DNA analysis at ~34,000BP. Additional pollen finds suggest the relative warm spell of the "Greenland interstadial (GI) 5" (two hundred kilometres east of Moscow, on the outskirts of Vladimir, near the Klyazma River). mtDNA analysis shows that the four individuals tested from Sungir belong to mtDNA Haplogroup U. The individual from Burial 1 belongs to mtDNA Haplogroup U8c, while the three individuals from Burial 2 belong to mtDNA Haplogroup U2. Y-DNA analysis shows that all four of the tested individuals from Sungir belong to Y-DNA Haplogroup C1a2.[32]

Kostenki-12Eastern Europe (Russia)Aurignacian33Dated at 32,600 ± 1,100 radio-carbon years, the find from Kostenki consists of a tibia and a fibula in a rich culture layer. The occupation layers contain bone and ivory artifacts, including possible figurative art, and fossil shells imported more than 500 kilometers.[33][34]
GowerWestern Europe (UK)Aurignacian33Despite its name, the Red Lady of Paviland is a partial skeleton of a young male, lacking a skull. It was discovered in 1823 in a cave burial in Gower, South Wales, United Kingdom. The bones were stained with ochre, and it was the first human fossil to have been found anywhere in the world. At 33,000 years old, it is still the oldest ceremonial burial of a modern human ever discovered anywhere in Western Europe.[35] Associated finds were red ochre anointing, a mammoth skull, and personal decorations suggesting shamanism or other religious practice. Grave goods are considered late Aurignacian or Early Gravettian in appearance. Genetic analysis of mtDNA yielded the Haplogroup H, the most common group in Europe.[36]
La Quina AvalWestern Europe (France)Aurignacian33Consisting of a partial juvenile mandible, the find is also associated with early Aurignacian tools. The jaw has some archaic features, though it is mainly modern.[37] The find is dated to max 33,000–32,000 radiocarbon years.[33]
Les Roisà MouthiersWestern Europe (France)Aurignacian32There are diagnostic modern human remains associated with a later Aurignacian assemblage at Les Roisà Mouthiers, France. The date is likely not older than 32,000 radiocarbon years.[33]
Mladeč cavesCentral Europe (Czech Republic)Aurignacian31The Mladeč caves in Moravia have yielded the remains of several individuals, but few artifacts. The artifacts found have tentatively been classified as Aurignacian. The finds have been radiocarbon dated to around 31,000 radiocarbon years (somewhat older in calendar years),[38] Mladeč 2 is dated to 31,320 +410, -390, Mladeč 9a to 31,500 +420, -400 and Mladeč 8 to 30,680 +380, -360 14C years.[33]
Peștera MuierilorSoutheastern Europe (Romania)Aurignacian31

The Peștera Muierilor ("Women's Cave") find is a single, fairly complete cranium of a woman with rugged facial traits and otherwise modern skull features, found in a lower gallery of the cave, among numerous cave bear remains. Radiocarbon dating yielded an age of 30,150 ± 800 years. No associated tools were found.[33]

Muierii and Cioclovina CavesSoutheastern Europe (Romania)Aurignacian30Cioclovina 1 is a complete neurocranium from a robust individual lacking all facial bones. The find is from a cave bear den, Cioclovina Cave, Romania. It is dated at 29,000 ± 700 radiocarbon years.[33][39]
Cro-MagnonWestern Europe (France)Aurignacian29
Cro-Magnon 2, the female skull found at Cro-Magnon (1884 drawing)

The original Cro-Magnon find was discovered in a rock shelter at Les Eyzies, Dordogne, France. The type specimen from the site is Cro-Magnon 1, a male, carbon dated to about 28,000 14C years old.[40] (27,680 ± 270 BP). Compared to Neanderthals, the skeletons showed the same high forehead, upright posture and slender (gracile) skeleton as modern humans. The other specimens from the site are a female, Cro-Magnon 2, and another male, Cro-Magnon 3. The condition and placement of the remains of Cro-Magnon 1, along with pieces of shell and animal teeth in what appear to have been pendants or necklaces, raises the question of whether they were buried intentionally. If Cro-Magnons buried their dead intentionally, it suggests they had a knowledge of ritual, by burying their dead with necklaces and tools, or an idea of disease and that the bodies needed to be contained.[41] Analysis of the pathology of the skeletons shows that the humans of this period led a physically difficult life. In addition to infection, several of the individuals found at the shelter had fused vertebrae in their necks, indicating traumatic injury; the adult female found at the shelter had survived for some time with a skull fracture. As these injuries would be life-threatening even today, this suggests that Cro-Magnons relied on community support and took care of each other's injuries.[41] The Abri of Cro-Magnon is part of the UNESCO World Heritage of the "Prehistoric Sites and Decorated Caves of the Vézère Valley".[42]

PředmostíCentral Europe (Czech Republic)Aurignacian-Gravettian26The Předmostí site, near Přerov, Moravia was discovered in the late 19th century. Excavations were conducted between 1884 and 1930. As the original material was lost during World War II, in the 1990s, new excavations were conducted.[43] The Předmostí site appears to have been a living area with associated burial ground with some 20 burials, including 15 complete human interments, and portions of five others, representing either disturbed or secondary burials. Cannibalism has been suggested to explain the apparent subsequent disturbance,[44] though it is not widely accepted. The non-human fossils are mostly mammoth. Many of the bones are heavily charred, indicating they were cooked. Other remains include fox, reindeer, ice-age horse, wolf, bear, wolverine, and hare. Remains of three dogs were also found, one of which had a mammoth bone in its mouth.[45]

The Předmostí site is dated to between 24,000 and 27,000 years old. The people were essentially similar to the French Cro-Magnon finds. Though undoubtedly modern, they had robust features indicative of a big-game hunter lifestyle. They also share square eye socket openings found in the French material.[19]

Balzi RossiItalyAurignacian-Gravettian25Grimaldi Man is a find from the Ligurian Coast in Italy. The caves yielded several finds, among them two fairly complete skeletons in the lower Aurignacian layer. Though the age and accompanying tools suggests Cro-Magnon, the skeletons differ physically from the large and robust Cro-Magnons, being slender and rather short.[46] The remains from one of the caves, the "Barma Grande", have in recent time been radiocarbon dated to 25 ka.[47]

The Venus figurines of Balzi Rossi have been dated to the later Gravettian or Epi-Gravettian, 24 to 19 ka.[48]

Abrigo do Lagar VelhoWestern Europe (Portugal)Gravettian24The Lapedo child from Abrigo do Lagar Velho, Portugal, about 24,000 years old, a fairly complete and quite robust skeleton, possibly showing some Neanderthal traits.[49]
Abri PataudWestern Europe (France)Solutrean21
Skull of the Pataud woman

The Abri Pataud shelter shows human habitation throughout the Aurignacian to Solutrean, but was abandoned in the early Magdalenian, about 17 ka. The "Pataud woman" is the name given to the remains of a young woman, about 20 years old, deposited with the body of a newborn child, and is about 21,000 years old. The woman's skull was buried separately, about four meters from the body, lying protected between stones.

ChanceladeWestern Europe (France)Magdalenian15
The Chancalade skull

Chancelade man, a short and stocky older man buried in Chancelade, France, was found with Magdalenian tools.[50] Several other more fragmentary finds, like the skeleton from Laugerie-Basse and the Duruthy cave near Sorde-l'Abbaye has traditionally been linked to the Chancelade man.[51] The morphological difference in the Chancelade skull compared to the "stockier" Cro-Magnon type has been taken as evidence for a Gravettian or Magdalenian-era influx of a different population unrelated to the Aurignacian EEMH.

Cap Blanc rock shelterWestern Europe (France)Magdalenian-Azilian14Magdalenian Girl is a female skeleton, discovered in Dordogne in 1911. Dating to the later Magdalenian, transitional to the beginning Mesolithic.
Grotte du BichonWestern Europe (Switzerland)Magdalenian-Azilian14Bichon man is the skeleton of a young male of the mesolithic hunter-gatherer (WHG) lineage.
Ripari VillabrunaItalyMagdalenian-Azilian14

Villabruna 1 is a skeleton, dated 14.1–13.8 ka, buried in a shallow pit, the head turned to the left with arms stretched touching the body, with grave goods typical of hunter-gatherer equipment. Villabruna 1 is the oldest bearer of Y-haplogroup R1b (R1b1a-L754* (xL389,V88)) found in Europe, and has been taken as a representative of the beginning post-LGM (Mesolithic) migration movement from the Near East.[24]

Genetics

The lineage of EEMH contributes substantially to the modern populations indigenous to Europe. EEMH populations during the Last Glacial Maximum were probably confined to refugia, associated with the Solutrean and Gravettian cultures. From there, Europe was re-peopled during the Holocene climatic optimum. The genetic contribution of these populations to modern Europeans is dubbed "West European Hunter-Gatherer" (WHG). The identification of the WHG component in modern populations is based on the analysis of the genome of a Mesolithic hunter-gatherer buried c. 8000 years ago in the Loschbour rock shelter in Müllerthal, near Heffingen, Luxembourg.[note 3]

According to Fu et al. (2016), the mesolithic WHG lineage (dubbed the "Villabruna Cluster") already contains post-LGM admixture from the Near East and Caucasus, ending the period of isolation in EEMH of c. 37 to 14 ka.[note 4]

The WHG populations mixed extensively with the expanding Early European Farmers (EEF) populations during the European Neolithic.[53]

In terms of unipaternal lineages, EEMH were descended from the patrilineal Y-DNA haplogroups Haplogroup IJ and C1,[note 5][5] and maternal mt-DNA haplogroup N (and descendant haplogroups R and U).[note 6] Y-haplogroup IJ likely arose still in Southwest Asia. haplogroup I emerged about 35 to 30 ka, either in Europe or West Asia. Y-haplogroup K2a* (K-M2308) is associated with Central Asia, found in Siberian Ust'-Ishim man, but also in the Proto-Aurignacian Oase 1 fossil (Romania). Mt-haplogroup U5 arose in Europe just prior to the LGM, between 35 and 25 ka.

"Bichon man", an Azilian (early Mesolithic) skeleton found in the Swiss Jura, was found to be associated with the WHG lineage. He was a bearer of Y-DNA haplogroup I2a and mtDNA haplogroup U5b1h.[23]

See also
  • Anatomically modern humans
  • Peopling of Eurasia
  • Neanderthal extinction – Extinction of Neanderthals around 40,000 years ago
  • Ancient North Eurasian – Archaeogenetic name for an ancestral genetic component
  • Genetic history of Europe

Notes
  1. The process leading to the development of smaller and more fine-boned humans seems to have begun at least 50,000–30,000 years ago.[6]
  2. "The term "Cro-Magnon" has no formal taxonomic status, since it refers neither to a species or subspecies nor to an archaeological phase or culture. The name is not commonly encountered in modern professional literature in English, since authors prefer to talk more generally of anatomically modern humans (AMH). They thus avoid a certain ambiguity in the label "Cro-Magnon", which is sometimes used to refer to all early moderns in Europe (as opposed to the preceding Neanderthals), and sometimes to refer to a specific human group that can be distinguished from other Upper Paleolithic humans in the region. Nevertheless, the term "Cro-Magnon" is still very commonly used in popular texts because it makes an obvious distinction with the Neanderthals, and also refers directly to people rather than to the complicated succession of archaeological phases that make up the Upper Paleolithic. This evident practical value has prevented archaeologists and human paleontologists from dispensing entirely with the idea of Cro-Magnons."[7]
  3. "We observe a striking contrast between Europe west of the Caucasus and the Near East in degree of relatedness to WHG. In Europe, there is a much higher degree of allele sharing with Loschbour man than with MA1, which we ascribe to the 60–80% WHG/(WHG+ANE) ratio in most Europeans that we report in SI14. In contrast, the Near East has no appreciable WHG ancestry but some ANE ancestry, especially in the northern Caucasus. (Jewish populations are marked with a square in this figure to assist in interpretation as their ancestry is often anomalous for their geographic regions.)"[52]
  4. "beginning with the Villabruna Cluster at least ~14,000 years ago, all European individuals analysed show an affinity to the Near East. This correlates in time to the Bølling-Allerød interstadial, the first significant warming period after the Glacial Maximum. Archaeologically, it correlates with cultural transitions within the Epigravettian in southern Europe and the Magdalenian-to-Azilian transition in western Europe. Thus, the appearance of the Villabruna Cluster may reflect migrations or population shifts within Europe at the end of the Ice Age, an observation that is also consistent with the evidence of mitochondrial DNA turnover".[24]
  5. Kostenki-14 (Russia): C1b, Goyet Q116-1 (Belgium) C1a.[24]
  6. Haplogroup N was found in two Gravettian-era fossils, Paglicci 52 Paglicci 12[11]

References
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